The orbicularis oris and buccinator muscles of mammals form an important subset of the facial musculature, the perioral muscles. In many taxa, these muscles form a robust muscular hydrostat capable of highly manipulative fine motor movements, likely accompanied by a specialized pattern of innervation. We conducted a retrograde nerve-tracing study of cranial nerve (CN) VII in pigs (Sus scrofa) to: (1) map the motor neuron pool distributions of the superior and inferior orbicularis oris, and the buccinator, to test the hypothesis that perioral muscle motor neuron pools exhibit a somatotopic organization within the facial motor nucleus; and (2) test the hypothesis that portions of the superior orbicularis oris (SOO) motor neuron pool also exhibit a somatotopic organization, reflecting a potential compartmentalization of function of the rostral, middle, and caudal segments of this muscle. Cresyl violet histological staining showed that the pig facial motor nucleus was comprised of 7 well-defined subnuclei. Neuroanatomical tracers injected into these perioral muscles transported to the motor neuron pools of the lateral 4 of the 7 subnuclei of the facial motor nucleus. The motor neuron pools of the perioral muscles were generally segregated from motoneurons innervating other facial muscles of the rostrum. However, motor neuron pools were not confined to single nuclei but instead spanned across 3–4 subnuclei. Perioral muscle motor neuron pools overlapped but were organized somatotopically. Motor neuron pools of portions of the SOO overlapped greatly with each other but exhibited a crude somatotopy within the SOO motor neuron pool. The large and somatotopically organized SOO motor neuron pool in pigs suggests that the upper lip might be more richly innervated than the other perioral muscles and functionally divided.

1.
Akintunde A, Buxton DF (1992) Quadruple labeling of brain-stem neurons: a multiple retrograde fluorescent tracer study of axonal collateralization. J Neurosci Methods 45:15–22.
2.
Ashwell KW (1982) The adult mouse facial nerve nucleus: morphology and musculotopic organization. J Anat 135:531–538.
3.
Baisden RH, Woodruff ML, Whittington DL, Baker DC, Benson AE (1987) Cells of origin of the branches of the facial nerve: A retrograde HRP study in the rabbit. Am J Anat 178:175–184.
4.
Boas JEV, Paulli S (1908) The Elephant’s Head. Jena: Gustav Fischer.
5.
Cacou C, Greenfield BE, Hunt NP, McGrouther DA (1996) Patterns of coordinated lower facial muscles function and their importance in facial reamination. Br J Plast Surg 49:274–280.
6.
Cantania KC, Kaas JH (1995) Organization of the somatosensory cortex of the Star-Nosed mole. J Comp Neurol 351:536–548.
7.
Cantania KC, Kaas JH (1997) Somatosensory fovea in the Star-Nosed mole: Behavioral use of the star in relation to innervation patterns and cortical representation. J Comp Neurol 387:215–233.
8.
Clancy B, Cauller LJ (1998) Reduction of background autofluorescence in brain sections following immersion in sodium borohydride. J Neurosci Methods 83:97–102.
9.
Clifford AB, Witmer LM (2004) Case studies in novel narial anatomy: 2. The enigmatic nose of the moose (Artiodactyla: Cervidae: Alces alces. J Zool (London) 262:339–360.
10.
Courville J (1966) The nucleus of the facial nerve: The relation between cellular groups and peripheral branches of the nerve. Brain Res 1:338–353.
11.
Dom RM (1982) Topographical representation of the peripheral nerve branches of the facial nucleus of the opossum: A study utilizing horseradish peroxidase. Brain Res 246:281–284.
12.
Dom RM, Falls W, Martin GF (1973) The motor nucleus of the facial nerve in the oppossum (Didelphis marsupialis virginiana). Its organization and connections. J Comp Neurol 152:373–402.
13.
Eisenberg JF (1981) The Mammalian Radiations: An Analysis of Trends in Evolution, Adaptation, and Behavior. Chicago: University of Chicago Press.
14.
Feinstein B, Lingegard E, Nyman E, Wohlfart G (1955) Morphologic studies of motor units in normal human muscles. Acta Anat 23:127–142.
15.
Freilinger G, Happak W, Burggasser G, Gruber H (1990) Histochemical mapping and fiber size analysis of facial muscles. Plast Reconst Surg 86:422–428.
16.
Frey R, Hofmann RR (1996) Evolutionary morphology of the proboscideal nose of Guenther’s dikdik (Rhynchotragus guentheri Thomas, 1894) (Mammalia, Bovidae). Zool Anz 235:31–51.
17.
Frey R, Hofmann RR (1996/97) Skull, proboscis musculature and preorbital gland in the saiga antelope and Guenther’s dikdik (Mammalia, Artiodactyla, Bovidae). Zool Anz 235:183–199.
18.
Friauf E, Herbert H (1985) Topographic organization of facial motor neurons to individual pinna muscles in rat (Rattus rattus) and bat (Rousettus aegyptiacus). J Comp Neurol 240:161–170.
19.
Getty R (1975) Ruminant myology. In Anatomy of Domestic Animals, Vol 1 (Grossman SA, ed). Philadelphia: WB Saunders Co., pp 791–797.
20.
Hannam AG, Wood WW, De Cou RE, Scott JD (1981) The effects of working side occlusal interferences on muscle activity and associated jaw movements in man. Arch Oral Biol 26:387–392.
21.
Hannam AG, McMillan AS (1994) Internal organization in the human jaw muscles. Crit Rev Oral Biol Med 5:55–89.
22.
Happak W, Burggasser G, Gruber H (1988) Histochemical characteristics of human mimic muscles. J Neurol Sci 83:25–35.
23.
Happak W, Burggasser JLG, Flowers A, Gruber H, Freilinger G (1997) Human facial muscles: dimensions, motor endplate distribution, and presence of muscle fibers with multiple motor endplates. Anat Rec 249:276–284.
24.
Herring SW (1972) The facial musculature of the Suoidea. J Morphol 137:49–62.
25.
Herring SW, Wineski LE, Anapol FC (1989) Neural organization of the masseter muscle in the pig. J Comp Neurol 280:563–576.
26.
Herring, SW (1992) Muscles of mastication: architecture and functional organization. In The Biological Mechanisms of Tooth Movement and Craniofacial Adaptation (Davidovitch Z, ed.). Columbus: Ohio State University Press, pp 541–548.
27.
Herring SW, Sola OM, Huang X, Zhang G, Hayashida N (1993) Compartmentalization in the pig latissimus dorsi muscle. Acta Anat 147:56–63.
28.
Hinrichsen CFL, Watson CD (1984) The facial nucleus of the rat: representation of facial muscles revealed by retrograde transport of horseradish peroxidase. Anat Rec 209:407–415.
29.
Hofmann RR (1989) Evolutionary steps of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive system. Oecologica 78:443–457.
30.
Horta-Júnior JAC, Tamega OJ, Cruz-Rizzolo RJ (2004) Cytoarchitecture and musculotopic organization of the facial motor nucleus in Cebus appella monkey. J Anat 204:175–190.
31.
Huang CS, Sirisko MA, Hiraba H, Murray GM, Sessle BJ (1988) Organization of the primate face motor cortex as revealed by intracortical microstimulation and electrophysiological identification of afferent inputs and corticobulbar projections. J Neurohysiol 59:796–818.
32.
Huber E (1930a) Evolution of facial musculature and cutaneous field of trigeminus. Part I. Quart Rev Biol 5:133–188.
33.
Huber E (1930b) Evolution of facial musculature and cutaneous field of trigeminus. Part II. Quart Rev Biol 5:389–437.
34.
Jacquin MF, Renehan WE, Rhoades, RW, Panneton WM (1993) Morphology and topography of identified primary afferents in trigeminal subnuclei principalis and oralis. J Neurophysiol 70:1911–1936.
35.
Jenny AB, Saper CB (1987) Organization of the facial nucleus and corticofacial projection in the monkey: a reconsideration of the upper motor neuron facial palsy. Neurology 37:930–939.
36.
Johnson JI (1990) Comparative development of somatic sensory cortex. In Cerebral Cortex: Comparative Structure and Evolution of Cerebral Cortex, Vol 8B, part II (Jones EG, Peters A, eds). New York: Plenum Press, pp 335–450.
37.
Katz AD, Catalano P (1987) The clinical significance of the various anastomotic branches of the facial nerve. Arch Otolaryngol Head Neck Surg 113:959–962.
38.
Kier WM, Smith KK (1985) Tongue, tentacles, and trunks: The biomechanics of movement in muscular hydrostats. Zool J Linn Soc 83:307–324.
39.
Klein BG, Rhoades RW (1985) Representation of whisker follicle intrinsic musculature in the facial motor nucleus of the rat. J Comp Neurol 232:55–69.
40.
Klein BG, Rhoades RW, Jacquin MF (1990) Topography of the facial musculature within the facial (VII) motor nucleus of the neonatal rat. Exp Brain Res 81:649–653.
41.
Komiyama M, Shibata H, Suzuki T (1984) Somatotopic representation of facial muscles within the facial nucleus of the mouse. Brain Behav Evol 24:1144–1151.
42.
Kume M, Uemura M, Matsude K, Matsushima R, Mizuno N (1978) Topographical representation of peripheral branches of the facial nerve within the facial nucleus: A HRP study in the cat. Neurosci Lett 8:5–8.
43.
Marshall CD, Huth GD, Edmonds VM, Halin DL, Reep RL (1998a) Prehensile use of perioral bristles during feeding and associated behaviors of the Florida manatee (Trichechus manatus latirostris). Marine Mamm Sci 14:274–289
44.
Marshall CD, Clark LA Reep RL (1998b) The muscular hydrostat of the Florida manatee (Trichechus manatus latirostris) and its role in the use of perioral bristles. Marine Mamm Sci 14:290–303.
45.
Marshall CD, Maeda H, Iwata M, Furuta M, Asano A, Rosas F, Reep RL (2003) Orofacial morphology and feeding behaviour of the dugong, Amazonian, West African and Antillean manatees (Mammalia: Sirenia): functional morphology of the muscular-vibrissal complex. J Zool (London) 259:1–16.
46.
Martin MR, Lodge D (1977) Morphology of the facial nucleus of the rat. Brain Res 123:1–12.
47.
Nickolai N (1953) Über die oberflächliche Facialismuskulatur des Schweines (Sus scrofa). Gegenbaurs Morphol Jahrbuch 93:321–363.
48.
Papez JW (1927) Subdivision of the facial nucleus. J Comp Neurol 43:159–191.
49.
Penfield W, Rasmussen T (1950) The Cerebral Cortex of Man. New York: MacMillan.
50.
Pokay MM (1991) The barrelettes – architectonic vibrissal representations in the brainstem trigeminal complex of the mouse. I. Normal structural organization. J Comp Neurol 309:161–199.
51.
Populin LC, Yin TC (1995) Topographical organization of the motoneuron pools that innervate the muscles of the pinna of the cat. J Comp Neurol 363:600–614.
52.
Provis J (1977) The organization of the facial nucleus of the brush-tailed possum (Trichosurus vulpecula). J Comp Neurol 172:177–188.
53.
Radpour S (1977) Organization of the facial nucleus in the cat. Laryngoscope 87:557–674.
54.
Radpour S, Gacek RR (1980) Facial nerve nucleus in the cat. Further study. Laryngoscope 90:685–692.
55.
Reighard J, Jennings HS (1935) Anatomy of the Cat. New York: Henry Holt and Co.
56.
Satoda T, Takahashi O, Tashiro T, Matsushima R, Uemura-Sumi M, Mizuno N (1987) Representation of the main branches of the facial nerve within the facial nucleus of the Japanese monkey (Macaca fuscata). Neurosci Lett 78:283–287.
57.
Satoda T, Takahashi O, Tashiro T, Matsushima R, Uemura-Sumi M, Mizuno N (1988) Somatotopic organization of facial nucleus of rabbit. With particular reference to intranuclear representation of perioral branches of the facial nerve. Anat Anz 165:83–90.
58.
Schaefer KP, Meyer DL, Schott D (1971) Optic and vestibular influences on ear movements. Brain Behav Evol 4:323–333.
59.
Schieber MH (2001) Constraints on somatotopic organization in the primary motor cortex. J Neurophysiol 86:2125–2143.
60.
Semba K, Egger MD (1986) The facial ‘motor’ nerve of the rat: control of vibrissal movement and examination of motor and sensory components. J Comp Neurol 247:144–158.
61.
Siegmund H, Santibañez-H G (1981) Effects of motor degeneration of the external ear muscles on the audio-visual targeting reflex in cats. Acta Neurobiol Exp 41:1–13.
62.
Siegmund H, Santibañez-H G (1982) Effector pattern of the audio-visual targeting reflex in cats. Acta Neurobiol Exp 42:311–326.
63.
Shigenaga Y, Okamoto T, Nishimori T, Suemune S, Nasution ID, Chen IC, Tsuru K, Yoshida A, Tabuchi K, Hosoi M, Tsuru H (1986a) Oral and facial representation in the trigeminal principal and rostral spinal nuclei of the cat. J Comp Neurol 244:1–18.
64.
Shigenaga Y, Suemune S, Nishimura M, Nishimori T, Sato H, Ishidori H, Yoshida A, Tsuru K, Tsuiki Y, Dateoka Y, Nasution ID, Hosoi M (1986b) Topographic representation of lower and upper teeth within the trigeminal sensory nuclei of adult cat as demonstrated by the transganglionic transport of horseradish peroxidase. J Comp Neurol 251:299–316.
65.
Stal P (1994) Characterization of human oro-facial and masticatory muscles with respect to fiber types, myosins and capillaries. Swed Dent J Suppl 98:1–55.
66.
Stern JT, Wells JP, Junger WL, Vangor AD, Fleagle JG (1980) An electromyographic study of the pectoralis major in the ateline and hylobates with special reference to the evolution of a pars clavicularis. Am J Phys Anthropol 52:12–25.
67.
Szentágothai J (1948) The representation of facial and scalp muscles in the facial nucleus. J Comp Neurol 88:207–220.
68.
Tsai TC, Wu CH, Wen CY, Shieh JY (1993) Studies of the motoneurons following the injection of horseradish peroxidase into the peripheral branches of the facial nerve in rats. Acta Anat 148:42–48.
69.
Uemura-Suma M, Manabee Y, Matsushima R, Mizuno N (1986) Correlation of the main peripheral branches of the facial nerve with the cytoarchitectonic subdivisions of the facial nucleus in the guinea pig. Anat Embryol 174:161–166.
70.
Van der Loos H (1976) Barreloids in mouse somatosensory thalamus. Neurosci Lett 2:1–6.
71.
Van Gehuchten A (1906) Anatomie du système nerveux de l’homme, 4th ed. Louvain: Uystpruyst-Dieudonné.
72.
Vrra-Jensen GF (1942) The Motor Nucleus of the Facial Nerve with a Survey of the Efferent Innervation of the Facial Muscles. Copenhagen: E. Munkagard.
73.
Watson CRR, Sakai S, Armstrong W (1982) Organization of the facial nucleus in the rat. Brain Behav Evol 20:19–28.
74.
Weijs, WA (1996) Functional somatotopic organization of motoneurons supplying the rabbit masseter muscle. J Comp Neurol 364:279–289.
75.
Weijs WA, Jüch PJW, Kwa SHS, Korfage JAM (1993) Motor unit territories and fiber types in rabbit masseter muscle. J Dent Res 72:1491–1498.
76.
Welker C (1971) Microelectrode delineation of fine grain somatotopic organization of SmI cerebral neocortex in albino rat. Brain Res 26:259–275.
77.
Welker W, Johnson JI, Noe A (1998) Major national resources for the study of brain anatomy. University of Wisconsin, Michigan State University and the National Museum of Health and Medicine. http://brainmuseum.org /index.html.
78.
Welt C, Abbs JH (1990) Musculotopic organization of the facial motor nucleus in Macaca fascicularis: A morphometric and retrograde tracing study with Cholera toxin B-HRP J Comp Neurol 291:621–636.
79.
Witmer LM, Sampson SD, Solounias N (1999) The proboscis of tapirs (Mammalia: Perissodactyla): a case study in novel narial anatomy. J Zool (London) 249:249–267.
80.
Wood WW (1986) A functional comparison of the deep and superficial parts of the human anterior temporal muscle. J Dent Res 65:924–926.
81.
Woolsey TA, Settlage PA, Meyer DR, Sencer W, Hamuy TP, Travis AM (1952) Patterns of localization in precentral and ‘supplementary’ motor areas and their relation to the concept of a premotor area. Res Pub Assoc Res Nerv Ment Dis 30:238–264.
82.
Woolsey TA, Van der Loos H (1970) The structural organization of layer IV in the somatosensory region (SI) of mouse cerebral cortex. The description of a cortical field composed of discrete cytoarchitectural units. Brain Res 17:205–242.
83.
Woolsey TA, Welker C, Schwartz RH (1975) Comparative anatomical studies of the SmI face cortex with special reference to the occurrence of ‘barrels’ in layer IV. J Comp Neurol 164:79–94.
84.
Yagita K (1910) Experimentelle Untersuchungen über den Urspung des Nervus facialis. Anat Anz 37:195–218.
85.
Yoshida A., Yasuda K, Dostrovsky JO, Bae YC, Takemura M, Shigenaga Y, Sessle BJ (1994) Two major types of premotoneurons in the feline trigeminal nucleus orals as demonstrated by intracellular staining with horseradish peroxidase. J Comp Neurol 347:4095–514.
Copyright / Drug Dosage / Disclaimer
Copyright: All rights reserved. No part of this publication may be translated into other languages, reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying, recording, microcopying, or by any information storage and retrieval system, without permission in writing from the publisher.
Drug Dosage: The authors and the publisher have exerted every effort to ensure that drug selection and dosage set forth in this text are in accord with current recommendations and practice at the time of publication. However, in view of ongoing research, changes in government regulations, and the constant flow of information relating to drug therapy and drug reactions, the reader is urged to check the package insert for each drug for any changes in indications and dosage and for added warnings and precautions. This is particularly important when the recommended agent is a new and/or infrequently employed drug.
Disclaimer: The statements, opinions and data contained in this publication are solely those of the individual authors and contributors and not of the publishers and the editor(s). The appearance of advertisements or/and product references in the publication is not a warranty, endorsement, or approval of the products or services advertised or of their effectiveness, quality or safety. The publisher and the editor(s) disclaim responsibility for any injury to persons or property resulting from any ideas, methods, instructions or products referred to in the content or advertisements.
You do not currently have access to this content.